The hemoglobins of the adult blackbird (Turdus merula, Passeriformes). The sequence of the major (HbA) and minor component (HbD)

The blood of the adult blackbird contains one major hemoglobin component (HbA = alpha A2, beta 2, ca. 80%) and one minor one (HbD = alpha D2 beta 2, ca. 20%). The Hb-components were separated by FPLC on a TSK SP-5 PW column, and eluted with a linear NaCl gradient, while the globin chains were purified on a cation exchange (CM-Cellulose). Tryptic peptides from the globin chains were separated by HPLC on an RP-2 Lichrosorb column. The complete amino acid sequence was determined by automatic Edman degradation, using film and gas phase methods. For the alpha A-, alpha D- and beta-chains, peptide alignment was carried out relative to the corresponding chains of the greylag goose (Anser anser). The close phylogenetic relationship between blackbird, tree sparrow and starling is verified by the hemoglobin sequence. The O2-affinities of the major and minor hemoglobin components of the blackbird are not yet known. Thus, the results were interpreted on the basis of primary structure. Substitutions of possible structural significance were examined with the help of molecular graphics/modelling.     

Interspecific Mate Choice Following Cross-fostering in a Mixed Colony of Greylag Geese (Anser anser) and Canada Geese (Branta canadensis). A Study on Development and Persistence of Species Preferences1

The role of early experience in mate choice and species preference of geese was studied in an experiment where young greylag geese (Anser anser) were cross-fostered by Canada geese (Branta canadensis). In two successive years, all eggs from 9 nests of wild Canada geese were removed and replaced by greylag goose eggs. The young greylag geese then followed their Canada goose foster parents on their southward migration in autumn, to return with them in spring. Of 35 returning birds, all 16 females paired with greylag goose males (100%) whereas of 19 males 5 paired with Canada goose females (14%) and the remaining 14 with greylag goose females (74%). The pair bonds generally persisted as long as both birds were present, but after loss of a partner, the remaining bird usually re-mated. Even when this happened several times during the lifetime of a male, the new mate was always a Canada goose female, showing that the males were sexually imprinted to this species. The Canada goose females which had mated with the greylag ganders also remated when widowed, but their new mate could be either a Canada goose or a greylag goose.     

The primary structure of the pallid bat (Antrozous pallidus, Chiroptera) hemoglobin

The complete primary structure of the hemoglobin from the Pallid Bat (Antrozous pallidus, Microchiroptera) is presented. This hemoglobin consists of two components with identical amino-acid sequences, differing, however, in the N-terminus which is formylated in 12.5% of the beta-chains. The alpha- and beta-chains were separated by reversed phase high performance liquid chromatography. The sequences of both chains were established by automatic Edman degradation with the film technique or gas phase method using the native chains and the tryptic peptides. The formylation of a part of the N-terminal peptide of the beta-chains was determined by mass spectrometric examination. Compared to the corresponding human chains we found 14 substitutions in the alpha-chains and 21 in the beta-chains. One substitution in the alpha-chains and three in the beta-chains are involved in alpha 1/beta 1-contacts. Among these the exchange beta 123(H1)Thr----Cys is unusual because cysteine was so far not found in this position of mammalian beta-chains. Compared to the hemoglobin of Myotis velifer, another representative of the family Vespertilionidae, 5 residues are replaced in the alpha-chains and 18 in the beta-chains.     

Isolation and characterization of microsatellite marker loci in the greylag goose (Anser anser)

Ten novel polymorphic microsatellite loci were isolated and characterized from the greylag goose, Anser anser, a long-term monogamous and biparental bird. Additionally, five new primers pairs were designed based on previously published microsatellite locus sequences from closely related species. Multiplex polymerase chain reactions conditions were optimized for all 15 primer pairs. The number of alleles ranged from two to 12 per locus with an observed heterozygosity ranging from 0.07 to 0.85. This marker set will be used to determine rates and origins of extra-pair and parasitic young in a population of individually banded greylag geese with known life histories.     

The cytogenetics of domestic geese

Hybrids were produced between an African male and several Pilgrim female domestic geese. Partial karyotypes revealed a difference in the fourth largest pair of autosomal chromosomes. This chromosome pair was metacentric in the African, submetacentric in the Pilgrim, and heteromorphic in the hybrids. A similar difference between the putative wild ancestors of the African and Pilgrim breeds has been reported by others. These findings provide cytological evidence to support the traditional opinion that the African breed was derived from the Asiatic swan goose (Anser cygnoides) and the Pilgrim breed was derived from the European greylag goose (Anser anser).     

Relationship between wild greylag and European domestic geese based on mitochondrial DNA

The origins of the European domestic goose are uncertain. The available information comes from archaeological findings and historical literature, but genetic evidence has hitherto been scarce. The domestic goose in Europe is derived from the greylag goose (Anser anser), but it is not known where the initial domestication took place and which of the two subspecies of greylag goose was ancestral. We aimed to determine the amount and geographical distribution of genetic diversity in modern populations of greylag geese as well as in different breeds of the domestic goose to make inferences about goose domestication. We studied DNA sequence variation in the mitochondrial control region of greylag geese from multiple populations across Europe and western Asia as well as specimens of domestic geese representing 18 modern breeds and individuals not belonging to any recognised breed. Our results show notable differences in genetic diversity between different greylag goose populations and the presence of six mitochondrial haplogroups which show a degree of geographical partitioning. The genetic diversity of the domestic goose is low, with 84% of sampled individuals having one of two major closely related haplotypes, suggesting that modern European domestic geese may derive from a narrow genetic base. The site of domestication remains unresolved, but domestic geese in Turkey were unusually diverse, indicating the importance of further sampling in the vicinity of the eastern Mediterranean and the Near East. There appears to be past or ongoing hybridisation between greylags and domestic geese in particular areas, consistent with field observations.     

Amino-acid sequences and functional differentiation of hemoglobins A and D from swift (Apus apus, Apodiformes)

The blood of the adult swift contains one major (HbA = alpha 2A beta 2) and two minor components (HbD = alpha 2D beta 2 and HbD'). The components were separated by FPLC with a TSK SP-5 PW-column in phosphate buffers, and were eluted with a linear NaCl gradient. HbD' could be detected only in freshly prepared hemolysates with the sensitive FPLC separation method. The globin chains were separated on a cation exchanger (CM-cellulose), the tryptic peptides by HPLC with a RP-2 LiChrosorb column. Their amino-acid sequences were determined by automatic Edman degradation with the film- or gas-phase method. For the alpha A-, alpha D- and beta-chains, peptide alignment was achieved by homologous comparison with the corresponding chains of the greylag goose (Anser anser). The structural significance of the substitutions was examined with the aid of molecular graphics. The oxygen-binding properties of the stripped hemolysate and of HbA and HbD and their dependence on pH, temperature and inositol polyphosphate are presented and discussed with reference to molecular structures and hypothermy that occurs during torpidity.     

Latitudinal-Related Variation in Wintering Population Trends of Greylag Geese (Anser Anser) along the Atlantic Flyway: A Response to Climate Change?

The unusually high quality of census data for large waterbirds in Europe facilitates the study of how population change varies across a broad geographical range and relates to global change. The wintering population of the greylag goose Anser anser in the Atlantic flyway spanning between Sweden and Spain has increased from 120 000 to 610 000 individuals over the past three decades, and expanded its wintering range northwards. Although population sizes recorded in January have increased in all seven countries in the wintering range, we found a pronounced northwards latitudinal effect in which the rate of increase is higher at greater latitudes, causing a constant shift in the centre of gravity for the spatial distribution of wintering geese. Local winter temperatures have a strong influence on goose numbers but in a manner that is also dependent on latitude, with the partial effect of temperature (while controlling for the increasing population trend between years) being negative at the south end and positive at the north end of the flyway. Contrary to assumptions in the literature, the expansion of crops exploited by greylag geese has made little contribution to the increases in population size. Only in one case (expansion of winter cereals in Denmark) did we find evidence of an effect of changing land use. The expanding and shifting greylag population is likely to have increasing impacts on habitats in northern Europe during the course of this century.     

The primary structure of the hemoglobin of the European marmot (Marmota marmota marmota, Rodentia)

The hemoglobin of the European marmot Marmota marmota marmota has been found to consist of only one component. In this work, we are presenting its primary structure. The globin chains have been separated by high performance liquid chromatography and the sequences have been determined by automated Edman degradation of the chains and their tryptic peptides, as well as of the peptide obtained by acid hydrolysis of the Asp-Pro bond in the beta-chains. In the alpha-chains we have found 13 and in the beta-chains 34 exchanges compared with the human alpha- and beta-chains, respectively. The amino acids which are substituted in the alpha-chains are not involved in any contacts, whereas in the beta-chains, one exchange involves a heme contact, two alpha 1/beta 1- and one alpha 1/beta 2-contacts. The functional and evolutionary aspects of these findings are discussed.     

Contributions of feather microstructure to eider down insulation properties

Insulation is an essential component of nest structure that helps provide incubation requirements for birds. Many species of waterfowl breed in high latitudes where rapid heat loss can necessitate a high energetic input from parents and use down feathers to line their nests. Common eider Somateria mollissima nest down has exceptional insulating properties but the microstructural mechanisms behind the feather properties have not been thoroughly examined. Here, we hypothesized that insulating properties of nest down are correlated to down feather (plumule) microstructure. We tested the thermal efficiency (fill power) and cohesion of plumules from nests of two Icelandic colonies of wild common eiders and compared them to properties of plumules of wild greylag goose Anser anser. We then used electron microscopy to examine the morphological basis of feather insulating properties. We found that greylag goose down has higher fill power (i.e. traps more air) but much lower cohesion (i.e. less prone to stick together) compared to common eider down. These differences were related to interspecific variation in feather microstructure. Down cohesion increased with the number of barbule microstructures (prongs) that create strong points of contact among feathers. Eider down feathers also had longer barbules than greylag goose down feathers, likely increasing their air‐trapping capacity. Feather properties of these two species might reflect the demands of their contrasting evolutionary history. In greylag goose, a temperate, terrestrial species, plumule microstructure may optimize heat trapping. In common eiders, a diving duck that nests in arctic and subarctic waters, plumule structure may have evolved to maximize cohesion over thermal insulation, which would both reduce buoyancy during their foraging dives and enable nest down to withstand strong arctic winds.     

Hemoglobin of tree sparrows (Passer montanus, Passeriformes): Sequence of the major (Hb A) and minor (Hb D) components

Blood of the adult Tree Sparrow (Passer montanus) contains two hemoglobin components, Hb A (ca. 85%), Hb D (ca. 15%). They differ in their alpha-chains (alpha A, alpha D), the beta-chains are identical. The complete primary structures of alpha A-, alpha D- and beta-chains are presented. Comparison with the Greylag Goose (Anser anser) hemoglobin (Hb A) showed that the alpha A-chains differ by 22 amino-acid exchanges, the beta-chains by 16. Comparison with the minor component of the Pheasant (Phasianus colchicus colchicus) hemoglobin (Hb D) showed that the alpha D-chains differ by 34 amino-acid exchanges. Proline is found incorporated in an internal position of an alpha-helix (pos. 124, H7). In comparison to that of the Starling (Sturnus vulgaris) the ratio of amino-acid exchanges for beta: alpha A: alpha D chains is 1 : 7 : 4; in comparison to other birds this ratio is found to be 1 : 2 (1.4-2.2):3 (2.2-4).     

The primary structure of the hemoglobin from the Australian ghost bat (Macroderma gigas, Microchiroptera).

The Australian ghost bat (Macroderma gigas, Microchiroptera) has two hemoglobin components in the ratio 3:2. They share identical beta-chains and differ by three replacements in the alpha-chains. The primary structures of all three chains are presented. They could be separated by high-performance liquid chromatography. The sequences were determined by automatic liquid and gas phase Edman degradation of the chains and their tryptic peptides. The two alpha-chains show 18 and 19 and the beta-chains 15 exchanges compared to human alpha- and beta-chains, respectively. The divergent evolution of Macroderma gigas and Megaderma lyra, two representatives of the family Megadermatidae, is discussed. An influence of replacements at functionally important positions on the hemoglobin oxygen affinity seems unlikely.     

Spatial and temporal feeding segregation of two Icelandic goose species during the spring pre-nesting period

The pre-nesting feeding behaviour of greylag Anser anser and pink-footed geese A brachyrhynchus was studied on agricultural land at low altitude in southern Iceland from 10 April to 8 May 1990 Greylag geese were already present on 12 April increased to 4580 birds by 24 April, but declined to 1300 by 3 May Pink-footed geese arrived around 20 April and numbers continued to increase to a peak count of 11340 on 3 May Over 60% of greylag geese initially used stubble fields on the coast where this habitat was most frequent, but increasingly resorted to grassland and wetland habitats during late April Later-arriving pink-feet predominantly used managed grassland, away from coastal areas At inland grassland sites, greylag numbers peaked on 20 April, pink-feet m early May The early exploitation by greylags was associated with grass growth initiated under protective snow-patches Greylags spent 90 times more time feeding within 1 m of snow patches with enhanced grass growth than expected by chance and their feeding rates near snow patches were faster and their step rates slower than further away By early May, grass growth was uniform and, although snow-patches persisted, no difference in forage quality, goose feeding rates or step rates could be detected It is concluded that, in spring 1990 at least, habitat segregation during spring migration in southern Iceland minimised competition between these two closely related goose species within the same geographical area In areas where both species exploit the same habitat, a two week difference m timing of breeding (and hence phenology of migration) further assures minimal overlap in feeding exploitation     

Carnivora: primary structure of the hemoglobins from ratel (Mellivora capensis)

The erythrocytes of adult ratel contain two hemoglobin components, with two alpha- and one beta-chains. In this paper, their complete amino acid sequences are presented. The two alpha-chains differ in one residue at position 34 (Ala----Val) only. The primary structure of the chains was determined by sequencing the N-terminal regions (45 steps) and the tryptic peptides after their isolation from the digests by reversed-phase high-performance liquid chromatography. The alignment of these peptides was deduced from homology with other carnivora globins. The alpha-chains show 21 and the beta-chains 11 exchanges compared with human globin chains. In the alpha-chains, one heme- and two alpha 1/beta 1 contacts are exchanged. In the beta-chains there are three exchanges which involve one alpha 1/beta 1-, one alpha 1/beta 2- and one heme-contact. Between the ratel hemoglobin and those of carnivora a high degree of homology was found.     

Carnivora: the primary structure of the Pacific Walrus (Odobenus rosmarus divergens, Pinnipedia) hemoglobin

The primary structure of the alpha- and beta-chains of the hemoglobin from the Pacific Walrus (Odobenus rosmarus divergens, Pinnipedia) is presented. Sequence analysis revealed only one hemoglobin component whereas two bands were found in polyacrylamide gel electrophoresis. The globin chains were separated by high-performance liquid chromatography and the sequences determined by automatic liquid- and gas-phase sequencing of the chains and their tryptic peptides. The alpha-chains show 20 and the beta-chains 12 exchanges compared to the corresponding human chains. In the alpha-chains one heme- and two alpha 1/beta 1-contacts were exchanged whereas in the beta-chains one alpha 1/beta 1-, one alpha 1/beta 2-and one heme-contact are substituted. Compared to Harbour Seal (Phoca vitulina) the Walrus hemoglobin shows 9 amino-acid replacements in the alpha-chains and 5 in the beta-chains. The relation between Pinnipedia and Arctoidea is discussed.     

The primary structure of the hemoglobin of an Indian flying fox (Cynopterus sphinx, Megachiroptera)

The hemoglobin of the Indian flying fox Cynopterus sphinx contains only one component. In this work, we are presenting its primary structure. The globin chains were separated by high-performance liquid chromatography and the sequences determined by automatic liquid and gas-phase Edman degradation of the chains and their tryptic peptides, as well as of the peptide obtained by acid hydrolysis of the Asp-Pro bond in the beta-chains. The alpha-chains show 14 and the beta-chains 19 exchanges compared with the human alpha- and beta-chains, respectively. In the alpha-chains one amino-acid exchange involves an alpha 1/beta 1 contact. In the beta-chains one heme contact, three alpha 1/beta 1- and one alpha 1/beta 2-contacts are exchanged. The functional and evolutionary aspects of these findings are discussed.     

The primary structure of pale-throated three-toed sloth (Bradypus tridactylus, Xenarthra) hemoglobin

The hemoglobin of the Pale-Throated Three-Toed Sloth (Bradypus tridactylus, Xenarthra) was separated into two components (ratio 4:1) with identical amino-acid analyses for the alpha- and beta-chains. The primary structures of both chains from the major component are given. They could be isolated by chromatography on carboxymethyl cellulose CM-52. The sequences have been determined by automatic Edman degradation of the native chains and their tryptic peptides. The comparison with human hemoglobin showed 27 substitutions in the alpha-chains and 33 in the beta-chains. In the alpha-chains one amino-acid exchange involves an alpha 1/beta 1-contact. In the beta-chains two heme- and four alpha 1/beta 1-contacts are substituted. The hemoglobin of the Sloth is compared to that of the Nine-Banded Armadillo (Dasypus novemcinctus), another representative of the order Xenerthra.     

The primary structure and functional properties of the hemoglobins of a ground squirrel (Spermophilus townsendii, Rodentia)

The hemoglobin of the ground squirrel Spermophilus townsendii consists of two components which are present in a ratio of ca. 2:1. The two hemoglobins have identical alpha-chains, but differ in their beta-chains. We present the primary structures of the alpha- and the two beta-globin chains. Following chain separation by chromatography on carboxymethyl-cellulose CM-52, the amino-acid sequences were established by automatic Edman degradation of the globin chains and the tryptic peptides, as well as of a peptide obtained by acid hydrolysis of the Asp-Pro bond of the beta-chains. The two beta-chains differ by only one amino-acid residue, Ala being present in the main and Asp in the minor component in position 58 (E2). The comparison with human hemoglobin showed only 14 exchanges in the alpha-chains but 33 in the beta-chains. Whereas no contact positions are affected in the alpha-chains, we found four such substitutions in the beta-chains, including one heme contact, two alpha 1/beta 1-contacts, and one alpha 1/beta 2-contact. It seems however, that the substitution found in the beta-chains has no effect on the oxygen affinity.     

Studies on monotreme proteins. VI. Amino acid sequence of the beta-chain of haemoglobin from the platypus, Ornithorhynchus anatinus.

The amino acid sequence of the 146 residues of the beta-chain of the major haemoglobin from the platypus has been determined. The soluble peptides derived from the chain by tryptic digestion were isolated by paper ionophoresis and chromatography. The amino acid sequences were determined by the dansyl-Edman procedure or by further digestion with other enzymes. The tryptic peptides were aligned by homology with other beta-globins. There were 14 changes in sequence compared with echidna beta-chain. The number of changes in sequence compared with human beta-chain is 34 which is less than the 39 changes between human and platypus alpha-chains. Generally there are more changes between beta-chains; there are only three other examples reported where there are more changes between alpha-chains than beta-chains, these are of echidna, rabbit and dog globins. By comparison with the 'contact sites' in horse haemoglobin there is one change in beta-haem contacts, three changes in beta1-alpha1 contacts and no changes in beta2-alpha1 contacts. The date of divergence of the monotremes from the other mammals was estimated at 132 +/- 33 million years, based on the number of amino acid differences between species and allowing for multiple mutations during the evolutionary period. This estimate differs widely from the estimate given by similar treatment of the alpha-chain sequences and the significance of this discrepancy to the validity of the method is discussed.     

Ontogenie und Funktion der Rangordnung innerhalb einer Geschwisterschar von Graugänsen (Anser anser L.)

Between the siblings of a family of greylag geese (Anser anser L.) a rank order is established by fighting during the first days after hatching, and also by other, more complex interactions. The position of each goose in the rank order can be defined by three groups of behaviour patterns (aggressive behaviour, “greetings”, escape responses). A clear-cut ranking relation between two individuals is a prerequisite for their close cohesion. The survival function of rank order lies in diminishing the intensity and frequency of agonistic interactions, in diminishing social stress situations and in guaranteeing the close family cohesion.